多寄主型寄生性天敌昆虫的寄主适应性及其影响因素

寄生性天敌昆虫对不同寄主资源的适应能力是其存活和繁殖的必要条件,它们的寄主选择行为则是其重要的适应结果。多寄主型寄生性天敌昆虫虽然对某种特定寄主资源的利用效率可能不如单寄主型寄生蜂,但却有利于拓展更宽的寄主范围,因此对环境的适应能力更强,更容易在自然界维持其种群的生存。统计结果也表明多寄主型天敌的生物防治效果往往比专食性天敌更高。有时生物防治成功的关键可能并不在于所利用的天敌种类的不同,而在于天敌的不同生物型或地理宗。总结了多寄主型寄生性天敌昆虫的寄主适应能力及其影响因素。寄生性天敌昆虫不仅因地理隔离产生种群分化,也可能因寄生不同的寄主产生种群内的分化,从而更加适应寄主的生活特性和栖境条件。寄主种类、寄主发育阶段、寄主大小、寄主营养、寄主免疫反应、寄主逃避反应、其它天敌的竞争、寄主共生或共栖生物的存在、寄主植物、天敌自身的学习能力及其共生微生物等多种因素对寄生性天敌昆虫的寄主适应性可产生影响。展望了多寄主型寄生性天敌昆虫对新寄主资源的拓展利用能力和适应性在生产上的可能应用前景和途径,以期为明确天敌与寄主间的互作关系,人工驯化寄生性天敌昆虫增强对靶标害虫的控制作用,合理利用天敌提高生物防治效率提供新的思路和理论支持。     

寄生蜂取食寄主特性及其在害虫生物防治中的作用

许多寄生性天敌昆虫的雌虫不仅寄生寄主, 而且还能取食寄主。在卵育型（synovigenic）寄生蜂类群中, 取食寄主行为是较为普遍的现象。本文综合近20年相关研究进展, 从寄生蜂类群、取食类型、生态学意义及影响因子等方面对寄生蜂的取食寄主行为进行了归纳总结。寄生蜂通过取食不仅可以杀死寄主, 直接起到控制害虫种群数量的作用, 还能通过取食策略为卵的成熟和再生提供营养来源, 对延长雌虫的寿命也有一定的帮助。对取食寄主行为的了解可为筛选优势寄生性天敌种类、评估寄生蜂在害虫生物防治中的作用提供重要信息。     

《昆虫的寄生性天敌和病原微生物》

全书分二卷。第一卷：寄生性天敌;第二卷：病原微生物。论述了昆虫与寄生性天敌和病原微生物的相互作用和影响。从基础生物学、生物化学和分子生物学的角度进行研究。第一卷：寄生性天敌（V．1Parasites）。共14章,364页。分1．昆虫寄生性天敌发育的模式;2寄生性天敌的作用：在寄生性天敌和寄主相互关系中蛋白质和肽所起的作用;3．昆虫内寄生性天敌和寄主相互关系中激素的作用;4．昆虫寄生性天敌和病原微生物引起的繁殖干扰;5．昆虫寄生性天敌和病原微生物对寄主行为的影响;6．寄生新陈代谢的变化对奇生性天敌营养的影响;7．畸形细…     

双斑截尾寄蝇对寄主种类及草地螟幼虫龄期和寄生部位的选择性

双斑截尾寄蝇Nemorilla maculosa Meigen是草地螟和其他鳞翅类昆虫的重要寄生天敌,但国内外对其寄生行为知之甚少。为了明确双斑截尾寄蝇的寄生行为特征及其对草地螟种群动态的影响,在22℃,光照16 h的条件下就其对不同寄主种类及草地螟幼虫龄期、寄主部位的选择性进行了研究。结果显示：在草地螟、甜菜夜蛾和粘虫3种幼虫同时存在的条件下,双斑截尾寄蝇对草地螟的寄生率显著高于甜菜夜蛾和粘虫,而对甜菜夜蛾的寄生率又显著高于粘虫。另外,幼虫被寄生的平均次数也呈现相同的趋势,表明双斑截尾寄蝇喜选择草地螟。在3, 4, 5龄草地螟幼虫同时存在时,双斑截尾寄蝇对5龄幼虫的寄生率和平均寄生次数均显著高于4龄和3龄幼虫的,表明该寄蝇主要选择5龄幼虫为其寄主。在被寄生的草地螟幼虫中,胸部被寄生的比率（66.4%）和次数（4.2）显著高于头部（23.4%,1.0）和腹部（10.9%,1.1）,而腹部及头部被寄生的比例及卵粒数没有显著差异,表明双斑截尾寄蝇主要选择幼虫的胸部进行寄生。最后,就双斑截尾寄蝇对产卵寄主选择的原因和机理进行了讨论。     

中国寄生蜂研究及其在害虫生物防治中的应用

寄生蜂是一类重要的寄生性天敌昆虫,种类繁多、习性复杂,在害虫生物防治和综合治理中发挥着极其重要的作用。在产卵时,寄生蜂携带的毒液、多DNA病毒等寄生因子就会随之进入寄主体内,发挥调控寄主生长、发育、免疫、代谢、行为的作用,从而保障了寄生蜂后代的发育。本文主要针对我国寄生蜂的系统分类、资源普查、生物学、生态学、寄主调控、人工繁殖、释放应用、田间保护和助增等方面的基础研究和应用进行了概述和整理。     

寄主植物-蚜虫-天敌三重营养关系的化学生态学研究进展

综述了寄主植物－蚜虫－天敌三重营养关系的化学生态学研究,重点阐述了3个研究热点：①植物挥发性物质在蚜虫及其天敌选择寄主行为过程中的作用;②蚜虫信息素和蜜露对蚜虫天敌寄主选择行为的影响;③植物挥发性物质对蚜虫信息系作用的影响。对寄主植物－蚜虫－天敌三重营养关系的全面了解,将为蚜虫的综合治疗提供新思维。     

皱胸尖裂姬蜂对机敏异漏斗蛛的拟寄生行为及其发育特征

寄生蜂是蜘蛛的主要天敌类群之一,但寄生于漏斗蛛的寄生蜂种类较少,且缺乏相关寄生蜂的详细研究报道。我们研究了皱胸尖裂姬蜂(Oxyrrhexis rugosus)对机敏异漏斗蛛(Allagelena difficilis)的拟寄生,旨在揭示蜘蛛寄生蜂的拟寄生行为及其发育特征。显微镜下观察了12头机敏异漏斗蛛头胸部背面的皱胸尖裂姬蜂卵,并以均值法统计了皱胸尖裂姬蜂各发育阶段的历期,进一步观察了皱胸尖裂姬蜂寄生机敏异漏斗蛛后对寄主的影响。交配后的雌性皱胸尖裂姬蜂经过寻找、降落、蛰刺蛛体后伺机将卵产在机敏异漏斗蛛的头胸部背面后部;孵化后的皱胸尖裂姬蜂幼虫头部形成一个特殊摄食导管,通过摄食导管获取蛛体的营养;幼虫随着龄期增长,体色由浅变深,历期约10 d;幼虫老熟后在蛛网的漏斗状管道内结茧化蛹,蛹期约为12 d;成体雌、雄蜂寿命约为11 d,皱胸尖裂姬蜂的平均生活史周期约为33 d。皱胸尖裂姬蜂寄生后对寄主蜘蛛的行为和生活状态造成了严重的影响,随着幼虫龄期增长被寄生机敏异漏斗蛛不再进食、蜕皮,活动减少,蛛体逐渐萎缩,直至老熟幼虫离开蛛体时,蜘蛛死亡。该研究不仅增加了漏斗蛛寄生性天敌的已知种类,而且初步了解了以漏斗蛛为寄主的姬蜂的产卵行为、后代的发育特征以及对寄主蜘蛛的影响,这将为研究蜘蛛寄生蜂及其与寄主蜘蛛之间的化学通讯和协同进化提供重要的基础资料。     

补充非寄主食物对芙新姬小蜂和潜蝇姬小蜂控害行为权衡的影响

芙新姬小蜂和潜蝇姬小蜂均为强卵育型(synovigenic)寄生蜂,成虫具有产卵寄生致死、寄主取食致死和寄主叮蛰直接致死3种寄主致死行为。为了明确成虫补充非寄主食物营养对2种姬小蜂控害潜力/控害行为的影响,本文比较研究了分别提供寄主食物和寄主食物+非寄主食物(10%葡萄糖)条件下的2种姬小蜂在初羽化前期(前6 d)内对美洲斑潜蝇幼虫的致死行为的影响。结果显示:在仅提供寄主食物时,芙新姬小蜂的寄生、取食和总体致死量均显著的高于潜蝇姬小蜂,但是直接致死能力却弱于后者;补充非寄主食物和仅提供寄主食物处理相比,两种寄生蜂的寄生量,取食量和总致死量均呈下降趋势。该研究显示:在初羽化前期,取食寄主食物的雌蜂比取食寄主食物+葡萄糖的雌蜂具有更强的致死能力。     

外寄生性花绒寄甲的寄生选择及其发育表现

"选择-表现"假说认为,成虫应该选择有利于子代发育的高品质寄主,但在寄主选择中,除了寄主品质外,其他因素也可能影响寄主选择决策。寄主选择研究通常以成虫为对象,而对那些初龄幼虫选择寄主的寄生性昆虫很少关注。以1龄幼虫积极搜寻寄主的寄生性花绒寄甲为模式生物,采用双选试验设计,观察了花绒寄甲初孵幼虫在不同体重青杨天牛幼虫之间、在已被寄生与健康的黄粉虫蛹之间的寄生选择性;然后采用回归设计,观察了花绒寄甲寄生若干不同体重的青杨天牛幼虫后的发育表现。研究结果表明,花绒寄甲1龄幼虫对体型较大的青杨天牛幼虫的选择偏好显著大于对体型较小的寄主幼虫的选择,选择大体型幼虫的比值比是选择小体型幼虫的4.55倍;对已被寄生的寄主黄粉虫蛹的选择偏好显著大于对健康寄主蛹的选择,选择已被寄生寄主的比值比是选择健康寄主的12.57倍。寄生青杨天牛幼虫的花绒寄甲幼虫发育历期平均为11.49 d、蛹历期为26.67 d、幼虫发育至成虫的羽化率50%,这些发育表现与寄生时青杨天牛幼虫的体重没有显著关系。但刚羽化寄甲成虫体重与寄生时寄主的体重存在显著的正直线关系:寄生时的寄主体重每增大0.01 g,羽化出的寄甲成虫体重增大近0.08%;方差分析寄甲成虫体重在不同寄主体重水平之间的差异表明,从体型较大寄主中羽化的寄甲成虫体重显著大于从体型较小寄主中羽化的成虫。研究结果说明,花绒寄甲初孵幼虫在寄主选择决策时,在寄主体型大小与被寄生状态之间可能采取折衷对策,而且对体型大小不同的寄主选择与子代发育适合度表现存在一致性,从而支持"选择-表现"假说。     

寄生蜂寻找隐蔽性寄主害虫的行为机制

林木蛀干类害虫具有高度的隐蔽性,是林业上的一类重要害虫,也是目前世界上最难防治的害虫类群之一.寄生蜂在与寄主长期的协同进化过程中,形成了搜索、发现和攻击寄主害虫的独特机制,能够有效地找到并寄生它们.总结了寄生性天敌寻找、发现并成功定位隐蔽性寄主害虫的行为学机制.寄生蜂可以利用来自嗅觉的化学信息物质(如寄主、寄主粪便、虫道共生菌的挥发性气味)、寄主成虫的化学通讯物质、来自视觉的植物表面色差信息、来自触觉的寄主保护物性状特征、来自寄主取食和运动所产生的介质振动信号以及来自寄主活动和代谢的红外辐射等多种途径有效地发现隐蔽性害虫的位置,从而完成寄生行为.有些寄生蜂还能综合利用来源不同的多种信息,从而提高寄主定位的可靠性和准确性.本文还对寄生蜂寻找寄主的这些线索在生物防治上可能的利用途径和前景进行了讨论.这对促进我国在该领域的研究,充分利用天敌昆虫,提高生物防治效率具有参考价值.     

寄生蜂寄主选择行为研究进展

寄生蜂是膜翅目昆虫中的重要类群,在自然界种类多,数量大。据文献报道全世界膜翅目昆虫大约有10万种”’,其中许多种类与农林害虫长期保持一定的制约关系,在害虫防治中占有重要地位。由于大量使用化学农药导致害虫的抗药性增加,利用天敌昆虫来防治害虫越来越显示出其重要性和优越性。但是,直到近20年来,寄生蜂的寄生行为才受到重视,并给予了广泛的、深入的研究”－‘’严格地讲,寄生蜂应该称为拟寄生蜂,它与通常所说的寄生昆虫像虱子、臭虫等有明显不同,主要区别在于：1．寄生仅仅发生在未成熟阶段,而成虫阶段营自由生活;2．寄…     

When parasitoids deal with the spatial distribution of their hosts: consequences for both partners

Insect parasitoids developing inside hosts face a true challenge: hosts are scattered in the field and their localization and selection require the use of complex and sometime confusing information. It was assumed for a long time that small-brained organisms like parasitoids have evolved simple and efficient behavioral mechanisms, leading them to be adapted to a given ecological situation, for example, the spatial distribution o f hosts in the habitat. However, hosts are not static and their distribution may also vary through generations and within the life of parasitoid individuals. We investigated if and how parasitoids deal with such a spatial com plexity in a m esocosm experiment. We used the Aphidius rhopalosiphi/Sitobion avenae parasitoid/host system to investigate if parasitoid females experiencing different host aggregation levels exhibit different foraging behaviors independently of the number of hosts in the environment. We showed that A. rhopalosiphi females exploited hosts more intensively both within and among patches at higher host aggregation levels. We discussed the adaptiveness of such behaviors in the light of evolution and biological control.     

Patch Time Allocation by the Parasitoid Diadegma semiclausum (Hymenoptera: Ichneumonidae). III. Effects of Kairomone Sources and Previous Parasitism

We investigated the effects of kairomone sources and previous parasitism on the patch-leaving behavior of Diadegma semiclausum, a solitary endoparasitoid of larval Plutella xylostella. Individual female wasps were released onto an experimental plant, and were allowed to freely leave for an alternative host plant placed upwind of the experimental plant in a wind tunnel. In one experiment, the experimental plant was either intact, contained host damage alone, or contained both hosts and host damage. In another experiment, the plant was infested with either unparasitized hosts, hosts parasitized previously by the female herself, or parasitized by Cotesia plutellae, another larval endoparasitoid of P. xylostella. We analyzed the influence of kairomone sources, host types, and within-patch foraging experience on the patch-leaving tendency of D. semiclausum by means of the proportional hazards model. Presence of host damage, and unsuccessful host encounters as a result of host defenses decreased the parasitoids' patch-leaving tendency, while successful oviposition, self-superparasitism, and rejection of parasitized hosts increased their patch-leaving tendency. A conceptual model of the parasitoid's patch-leaving behavior is proposed on the basis of the results of current and previous studies.     

Learning by the parasitoid wasp, Aphidius ervi (Hymenoptera: Braconidae), alters individual fixed preferences for pea aphid color morphs

Learning, defined as changes in behavior that occur due to past experience, has been well documented for nearly 20 species of hymenopterous parasitoids. Few studies, however, have explored the influence of learning on population-level patterns of host use by parasitoids in field populations. Our study explores learning in the parasitoid Aphidius ervi Haliday that attacks pea aphids, Acyrthosiphon pisum (Harris). We used a sequence of laboratory experiments to investigate whether there is a learned component in the selection of red or green aphid color morphs. We then used the results of these experiments to parameterize a model that examines whether learned behaviors can explain the changes in the rates of parasitism observed in field populations in South-central Wisconsin, USA. In the first of two experiments, we measured the sequence of host choice by A. ervi on pea aphid color morphs and analyzed this sequence for patterns in biased host selection. Parasitoids exhibited an inherent preference for green aphid morphs, but this preference was malleable; initial encounters with red aphids led to a greater chance of subsequent orientation towards red aphids than predicted by chance. In a second experiment, we found no evidence that parasitoids specialize on red or green morphs; for the same parasitoids tested in trials separated by 2 h, color preference in the first trial did not predict color preference in the second, as would be expected if they differed in fixed preferences or exhibited long-term (> 2 h) learning. Using data from the two experiments, we parameterized a population dynamics model and found that learning of the magnitude observed in our experiments leads to biased parasitism towards the most common color morph. This bias is sufficient to explain changes in the ratio of aphid color morphs observed in field sites over multiple years. Our study suggests that for even relatively simple organisms, learned behaviors may be important for explaining the population dynamics of their hosts.     

Patch Time Allocation by the Parasitoid <Emphasis Type="Italic">Diadegma semiclausum</Emphasis> (Hymenoptera: Ichneumonidae). II. Effects of Host Density and Distribution

This study investigated the effects of host density and distribution on the patch-leaving behavior of Diadegma semiclausum (Hymenoptera: Ichneumonidae), a solitary endoparasitoid of larval Plutella xylostella (Lepidoptera: Plutellidae). Individual female wasps were released onto an experimental plant infested with host larvae at different densities and distributions, and were allowed to freely leave for an alternative host plant placed upwind of the experimental plant in a wind tunnel. The influence of host density and distribution, as well as within-patch foraging experience, on the parasitoids patch-leaving tendency was analyzed by means of the proportional hazards model. This study aimed to test the predictions of a number of patch-leaving models, including the Marginal Value Theorem, rules of thumb, and incremental or countdown mechanisms. The parasitoids patch-leaving tendency decreased with increased host density, more clustered host distribution, and unsuccessful host encounter as a result of host defense, but increased with successful oviposition. None of the simple rules of thumb such as fixed time, fixed number of hosts parasitized, or fixed giving-up time was employed by this parasitoid. The results agreed with the general predictions of the Marginal Value Theorem that patch residence time and numbers of ovipositions by the parasitoid increased with increasing host density. The decreasing influence of oviposition on the parasitoids patch-leaving tendency, regardless of host density or distribution, was consistent with the prediction of a countdown mechanism.     

Genetic variation in patch time allocation in a parasitic wasp

1. The intra-patch experience acquired by foraging parasitoid females has often been considered to have a strong influence on their tendency to leave a patch, and thus on their total patch residence time. Most studies that have been performed on this subject suggest that the patch-leaving rules observed are adaptive because they enable the females to adjust their patch residence time to local environmental conditions.
2. Considering a behavioural rule as being adaptive supposes that it has been progressively settled by natural selection, and thus that there is, in the population, genetic variation on which the natural selection could act.
3. Therefore, this study aimed to discover whether there was indeed genetic variability in the patch-leaving decision rules in a population of the egg parasitoid species Telenomus busseolae , which attacks patches of its hosts, the eggs of Sesamia nonagrioides . Different wasp families were compared using the isofemale lines method, and the behavioural records were analysed by means of a modified version of the Cox's proportional hazards model proposed by Haccou et al . (1991 ) and Hemerik, Driessen & Haccou (1993 ).
4. The results obtained show that T. busseolae females increase their tendency to leave the patch after each successful oviposition. Each host rejection also led to an increase in the tendency to leave the patch, but this effect was smaller when host rejections were observed between two ovipositions occurring in rapid succession. Subsequent visits to the patch also increased the patch-leaving tendency.
5. Genetic variability was found in both the global patch-leaving tendency and in the effect that successful ovipositions and host rejections have on this tendency.
6. The adaptive and evolutionary consequences of these results are discussed.     

Influence of environmental conditions on patch exploitation strategies of parasitoids

In patchy environments, patch-leaving decision rules are a keycomponent of the foraging behavior of parasitoids that haveto share their searching time between the different host patchesto optimize the number of ovipositions. It has been experimentallyshown that one of the proximate cues involved in patch-leavingdecision is the number of ovipositions performed by the parasitoidwhile in the patch. Ovipositions can have either a decrementalor an incremental effect on the patch residence time dependingon parasitoid species. Previous theoretical studies have suggestedthat environmental conditions and the ability of parasitoidsto reliably estimate the number of available hosts on a patchwhen entering it should influence how patch departure dependson ovipositions. We developed an individual-based model to testa large variety of patch-leaving decision rules in differentenvironmental conditions. This model includes a more realisticrepresentation of individual acquisition of information thanprevious theoretical work. In accordance with previous results,we show that the best use of information depends on environmentalconditions. Moreover, we identify the environmental factorsthat have a decisive influence on the most appropriate typeof rule (incremental or decremental). Decremental mechanismsare most efficient only in restricted conditions characterizedby a large number of patches and few parasitoids. The sensitivityof decision rules to frequency-dependent processes was testedby numerical invasibility experiments. Rare mutants using anyalternative rule never outperform populations using a high-performancerule. Incremental rules seem to be more sensitive than decrementalones to frequency-dependent processes.     

Parasitic wasps learn and report diverse chemicals with unique conditionable behaviors

Parasitoids exploit numerous chemical cues to locate hosts and food. Whether they detect and learn chemicals foreign to their natural history has not been explored. We show that the parasitoid Microplitis croceipes can associate, with food or hosts, widely different chemicals outside their natural foraging encounters. When learned chemicals are subsequently detected, this parasitoid manifests distinct behaviors characteristic with expectations of food or host, commensurate with prior training. This flexibility of parasitoids to rapidly link diverse chemicals to resource needs and subsequently report them with recognizable behaviors offers new insights into their foraging adaptability, and provides a model for further dissection of olfactory learning related processes.     

DEFENSE TRAITS OF LARVAL DROSOPHILA MELANOGASTER EXHIBIT GENETICALLY BASED TRADE‐OFFS AGAINST DIFFERENT SPECIES OF PARASITOIDS

Populations of Drosophila melanogaster face significant mortality risks from parasitoid wasps that use species‐specific strategies to locate and survive in hosts. We tested the hypothesis that parasitoids with different strategies select for alternative host defense characteristics and in doing so contribute to the maintenance of fitness variation and produce trade‐offs among traits. We characterized defense traits of Drosophila when exposed to parasitoids with different host searching behaviors (Aphaereta sp. and Leptopilina boulardi). We used host larvae with different natural alleles of the gene Dopa decarboxylase (Ddc), a gene controlling the production of dopamine and known to influence the immune response against parasitoids. Previous population genetic analyses indicate that our focal alleles are maintained by balancing selection. Genotypes exhibited a trade‐off between the immune response against Aphaereta sp. and the ability to avoid parasitism by L. boulardi. We also identified a trade‐off between the ability to avoid parasitism by L. boulardi and larval competitive ability as indicated by differences in foraging and feeding behavior. Genotypes differed in dopamine levels potentially explaining variation in these traits. Our results highlight the potential role of parasitoid biodiversity on host fitness variation and implicate Ddc as an antagonistic pleiotropic locus influencing larval fitness traits.